Abstract: Great ape gesture has become a research topic of intense interest, because its intentionality and flexibility suggest strong parallels to human communication. Yet the fundamental question of whether an animal species’ gestures carry specific meanings has hardly been addressed. We set out a systematic approach to studying intentional meaning in the gestural communication of non-humans and apply it to a sample of orangutan gestures. We propose that analysis of meaning should be limited to gestures for which (1) there is strong evidence for intentional production and (2) the recipient’s final reaction matches the presumed goal of the signaller, as determined independently. This produces a set of successful instances of gesture use, which we describe as having goal–outcome matches. In this study, 28 orangutans in three European zoos were observed for 9 months. We distinguished 64 gestures on structural grounds, 40 of which had frequent goal–outcome matches and could therefore be analysed for meaning. These 40 gestures were used predictably to achieve one of 6 social goals: to initiate an affiliative interaction (contact, grooming, or play), request objects, share objects, instigate co-locomotion, cause the partner to move back, or stop an action. Twenty-nine of these gestures were used consistently with a single meaning. We tested our analysis of gesture meaning by examining what gesturers did when the response to their gesture did not match the gesture’s meaning. Subsequent actions of the gesturer were consistent with our assignments of meaning to gestures. We suggest that, despite their contextual flexibility, orangutan gestures are made with the expectation of specific behavioural responses and thus have intentional meanings as well as functional consequences.

Abstract: Longitudinal behavioral data generally contains a significant amount of structure. In this work, we identify the structure inherent in daily behavior with models that can accurately analyze, predict, and cluster multimodal data from individuals and communities within the social network of a population. We represent this behavioral structure by the principal components of the complete behavioral dataset, a set of characteristic vectors we have termed eigenbehaviors. In our model, an individual’s behavior over a specific day can be approximated by a weighted sum of his or her primary eigenbehaviors. When these weights are calculated halfway through a day, they can be used to predict the day’s remaining behaviors with 79% accuracy for our test subjects. Additionally, we demonstrate the potential for this dimensionality reduction technique to infer community affiliations within the subjects’ social network by clustering individuals into a behavior space spanned by a set of their aggregate eigenbehaviors. These behavior spaces make it possible to determine the behavioral similarity between both individuals and groups, enabling 96% classification accuracy of community affiliations within the population-level social network. Additionally, the distance between individuals in the behavior space can be used as an estimate for relational ties such as friendship, suggesting strong behavioral homophily amongst the subjects. This approach capitalizes on the large amount of rich data previously captured during the Reality Mining study from mobile phones continuously logging location, proximate phones, and communication of 100 subjects at MIT over the course of 9 months. As wearable sensors continue to generate these types of rich, longitudinal datasets, dimensionality reduction techniques such as eigenbehaviors will play an increasingly important role in behavioral research.

Abstract: Coordination of primate group movements by individual group members is generally categorized as leadership behavior, which entails several steps: deciding where to move next, initiating travel, and leading a group between food, water sources, and rest sites. Presumably, leaders are able to influence their daily foraging efficiency and nutritional intake, which could influence an individual's feeding ecology and long-term reproductive success. Within anthropoid species, females lead group movements in most female-bonded groups, while males lead groups in most nonfemale-bonded groups. Group leadership has not been described for social prosimians, which are typically not female-bonded. We describe group movements in two nonfemale-bonded, lemurid species living in southeastern Madagascar, Propithecus diadema edwardsi and Eulemur fulvus rufus. Although several social lemurids exhibit female dominance Eulemur fulvus rufus does not, and evidence for female dominance is equivocal in Propithecus diadema edwardsi. Given the ecological stresses that females face during reproduction, we predict that females in these two species will implement alternative behavioral strategies such as group leadership in conjunction with, or in the absence of, dominance interactions to improve access to food. We found that females in both species initiated and led group movements significantly more often than males did. In groups with multiple females, one female was primarily responsible for initiating and leading group movements. We conclude that female nutritional needs may determine ranging behavior to a large extent in these prosimian species, at least during months of gestation and lactation.

Abstract: Animal groups arise from individuals’ choices about the number, characteristics, and identity of associates. Individuals make these choices to gain benefits from their associations. As the needs of an individual change with its phenotype, so too we expect the nature of its associations to vary. In this paper, we investigate how the social priorities of male plains zebra (Equus burchelli) depend on reproductive state. An adult male is either a bachelor, and lacking mating access, or a stallion defending a harem. Multiple harems and bachelor males aggregate in larger herds. Herds frequently split and merge, affording males opportunities to change associates. Over a 4-year period, we sampled the herd associations in a population of 500–700 zebras. To isolate the effects of reproductive state on male social behavior, we account for potential confounding factors: changes in population size, grouping tendencies, and sampling intensity. We develop a generally applicable permutation procedure, which allows us to test the null hypothesis that social behavior is independent of male status. Averaging over all individuals in the population, we find that a typical bachelor is found in herds containing significantly more adults, bachelors, and stallions than the herds of a typical stallion. Further, bachelors’ bonds with each other are more persistent over time than those among stallions. These results suggest that bachelors form cohesive cliques, in which we may expect cooperative behaviors to develop. Stallion–stallion associations are more diffuse, and less conducive to long-term cooperation.

Abstract: Researchers are increasingly turning to network theory to understand the social nature of animal populations. We present a computational framework that is the first step in a series of works that will allow us to develop a quantitative methodology of social network sampling to aid ecologists in their social network data collection. To develop our methodology, we need to be able to generate networks from which to sample. Ideally, we need to perform a systematic study of sampling protocols on different known network structures, as network structure might affect the robustness of any particular sampling methodology. Thus, we present a computational tool for generating network structures that have user-defined distributions for network properties and for key measures of interest to ecologists. The user defines the values of these measures and the tool will generate appropriate network randomizations with those properties. This tool will be used as a framework for developing a sampling methodology, although we do not present a full methodology here. We describe the method used by the tool, demonstrate its effectiveness, and discuss how the tool can now be utilized. We provide a proof-of-concept example (using the assortativity measure) of how such networks can be used, along with a simulated egocentric sampling regime, to test the level of equivalence of the sampled network to the actual network.

Abstract: While viewing faces, human adults often demonstrate a natural gaze bias towards the left visual field, that is, the right side of the viewee’s face is often inspected first and for longer periods. Using a preferential looking paradigm, we demonstrate that this bias is neither uniquely human nor limited to primates, and provide evidence to help elucidate its biological function within a broader social cognitive framework. We observed that 6-month-old infants showed a wider tendency for left gaze preference towards objects and faces of different species and orientation, while in adults the bias appears only towards upright human faces. Rhesus monkeys showed a left gaze bias towards upright human and monkey faces, but not towards inverted faces. Domestic dogs, however, only demonstrated a left gaze bias towards human faces, but not towards monkey or dog faces, nor to inanimate object images. Our findings suggest that face- and species-sensitive gaze asymmetry is more widespread in the animal kingdom than previously recognised, is not constrained by attentional or scanning bias, and could be shaped by experience to develop adaptive behavioural significance.

Abstract: Matrilineal kin-relations among wild bonobos (Pan paniscus) were studied by DNA analysis. Subject individuals were the members of E1 group, living at Wamba, Zaire, which has been studied since 1974. DNA samples were extracted from wadges that bonobos spat out when feeding on sugar cane. The D-loop region of mitochondrial DNA was amplified by the PCR method, and a nucleotide sequence of 350 base pairs was determined for 17 individuals. Nucleotide variations were found at 44 positions of the sequence. Based on these variations, 13 matrilineal units were divided into seven groups, and the mother of an orphan male was determined among several females. These genetic analyses, together with behavioral observation to date, revealed the following facts. High sequence variation in the target region indicated that females transfer between groups of bonobos, which is in agreement with supposition from long-term field studies. For females, there was no relationship between genetic closeness and social closeness that is represented by frequencies of proximity or grooming. After immigration into a new group, females form social associations with senior females without regard to kin relationship.

Abstract: Place learning abilities represent adaptations that contribute also to foraging efficiency under given spatio-temporal conditions. We investigated if this ability in turn constrains decision making in two sympatric vole species: while the herbivorous common vole (Microtus arvalis) feeds on spatio-temporally predictable food resources (e.g. roots, tubers and shoots of plant tubers), the omnivorous bank vole (Myodes glareolus) additionally subsists on temporally unpredictable food resources (e.g. insects and seeds). Here, we compare the spatial reference memory and working memory of the two species. In an automated operant home cage with eight water places, female voles either had to learn the fixed position of non-depletable places (reference memory task) or learn and avoid previously visited water places depleted in a single visit (win-shift task). In the reference memory task, Microtus females required significantly more choices to find all water places, initially performed slightly worse than Myodes females, and displayed slightly lower asymptotic performance. Both species were highly similar in new learning of the same task. In the more complex win-shift task, asymptotic performance was significantly lower in Microtus (72% correct) than in Myodes (79%). Our results suggest that both vole species resemble each other in their efficiency to exploit habitats with low spatio-temporal complexity but may differ in their efficiency at exploiting habitats with temporally changing spatial food distributions. The results imply that spatial ability adjusted to specific food distributions may impair flexible use of habitats that differ in their food distribution and therefore, decrease a species’ chances of survival in highly dynamic environments.

Abstract: There is an established and very influential view that primate societies have identifiable, persistent social organizations. It assumes that association patterns reflect long-term strategic interests that are not qualitatively perturbed by short-term environmental variability. We used data from two baboon troops in markedly different habitats over three consecutive seasons to test this assumption. Our results demonstrate pronounced cyclicity in the extent to which females maintained differentiated relationships. When food was plentiful, the companionships identified by social network analysis in the food-scarce season disappeared and were replaced by casual acquaintanceships more representative of mere gregariousness. Data from the fourth, food-scarce, season at one site indicated that few companions were re-united. It is likely that this reflected stochastic variation in individual circumstances. These results suggest that attention could profitably be paid to the effects of short-term local contingencies on social dynamics, and has implications for current theories of primate cognitive evolution.

Abstract: Great ape gestural communication is known to be intentional, elaborate and flexible; yet there is controversy over the best interpretation of the system and how gestures are acquired, perhaps because most studies have been made in restricted, captive settings. Here, we report the first systematic analysis of gesture in a population of wild chimpanzees. Over 266 days of observation, we recorded 4,397 cases of intentional gesture use in the Sonso community, Budongo, Uganda. We describe 66 distinct gesture types: this estimate appears close to asymptote, and the Sonso repertoire includes most gestures described informally at other sites. Differences in repertoire were noted between individuals and age classes, but in both cases, the measured repertoire size was predicted by the time subjects were observed gesturing. No idiosyncratic usages were found, i.e. no gesture type was used only by one individual. No support was found for the idea that gestures are acquired by ‘ontogenetic ritualization’ from originally effective actions; moreover, in detailed analyses of two gestures, action elements composing the gestures did not closely match those of the presumed original actions. Rather, chimpanzee gestures are species-typical; indeed, many are ‘family-typical’, because gesture types recorded in gorillas, orangutans and chimpanzee overlap extensively, with 24 gestures recorded in all three genera. Nevertheless, chimpanzee gestures are used flexibly across a range of contexts and show clear adjustment to audience (e.g. silent gestures for attentive targets, contact gestures for inattentive ones). Such highly intentional use of a species-typical repertoire raises intriguing questions for the evolution of advanced communication.